Sustainability is a key principle in natural resource management, and it involves operational efficiency, minimisation of environmental impact and socio-economic considerations; all of which are interdependent. It has become increasingly obvious that continued reliance on fossil fuel energy resources is unsustainable, owing to both depleting world reserves and the green house gas emissions associated with their use. Therefore, there are vigorous research initiatives aimed at developing alternative renewable and potentially carbon neutral solid, liquid and gaseous biofuels as alternative energy resources. However, alternate energy resources akin to first generation biofuels derived from terrestrial crops such as sugarcane, sugar beet, maize and rapeseed place an enormous strain on world food markets, contribute to water shortages and precipitate the destruction of the world's forests. Second generation biofuels derived from lignocellulosic agriculture and forest residues and from non-food crop feedstocks address some of the above problems; however there is concern over competing land use or required land use changes. Therefore, based on current knowledge and technology projections, third generation biofuels specifically derived from microalgae are considered to be a technically viable alternative energy resource that is devoid of the major drawbacks associated with first and second generation biofuels. Microalgae are photosynthetic microorganisms with simple growing requirements (light, sugars, CO 2 , N, P, and K) that can produce lipids, proteins and carbohydrates in large amounts over short periods of time. These products can be processed into both biofuels and valuable co-products. This study reviewed the technologies underpinning microalgae-to-biofuels systems, focusing on the biomass production, harvesting, conversion technologies, and the extraction of useful co-products. It also reviewed the synergistic coupling of microalgae propagation with carbon sequestration and wastewater treatment potential for mitigation of environmental impacts associated with energy conversion and utilisation. It was found that, whereas there are outstanding issues related to photosynthetic efficiencies and biomass output, microalgae-derived biofuels could progressively substitute a significant proportion of the fossil fuels required to meet the growing energy demand.

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Biofuels from microalgae—A review of technologies for production, processing, and extractions of biofuels and co-products

Examinination of microbial diversity in environments of increasing salt concentrations indicates that certain types of dissimilatory metabolism do not occur at the highest salinities. Examples are methanogenesis for H2 + CO2 or from acetate, dissimilatory sulfate reduction with oxidation of acetate, and autotrophic nitrification. Occurrence of the different metabolic types is correlated with the free-energy change associated with the dissimilatory reactions. Life at high salt concentrations is energetically expensive. Most bacteria and also the methanogenic archaea produce high intracellular concentrations of organic osmotic solutes at a high energetic cost. All halophilic microorganisms expend large amounts of energy to maintain steep gradients of NA+ and K+ concentrations across their cytoplasmic membrane. The energetic cost of salt adaptation probably dictates what types of metabolism can support life at the highest salt concentrations. Use of KCl as an intracellular solute, while requiring far-reaching adaptations of the intracellular machinery, is energetically more favorable than production of organic-compatible solutes. This may explain why the anaerobic halophilic fermentative bacteria (order Haloanaerobiales) use this strategy and also why halophilic homoacetogenic bacteria that produce acetate from H2 + CO2 exist whereas methanogens that use the same substrates in a reaction with a similar free-energy yield do not.

Bioenergetic Aspects of Halophilism

The removal of nitrogen (N) in aquatic ecosystems is of great interest because excessive nitrate in groundwater and surface water is a growing problem. High nitrate loading degrades water quality and is linked to eutrophication and harmful algal blooms, especially in coastal marine waters. Past research on nitrate removal processes has emphasized plant or microbial uptake (assimilation) or respiratory denitrification by bacteria. The increasing application of stable isotopes and other tracer techniques to the study of nitrate removal has yielded a growing body of evidence for alternative, microbially mediated processes of nitrate transformation. These include dissimilatory (the reduction of nitrogen into other inorganic compounds, coupled to energy producing processes) reduction of nitrate to ammonium (DNRA), chemoautotrophic denitrification via sulfur or iron oxidation, and anaerobic ammonium oxidation (anammox), as well as abiotic nitrate removal processes. Here, we review evidence for the importance of...

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Have we overemphasized the role of denitrification in aquatic ecosystems? A review of nitrate removal pathways

. Water masses can become undersaturated with oxygen when natural processes alone or in combination with anthropogenic processes produce enough organic carbon that is aerobically decomposed faster than the rate of oxygen re-aeration. The dominant natural processes usually involved are photosynthetic carbon production and microbial respiration. The re-supply rate is indirectly related to its isolation from the surface layer. Hypoxic water masses ( Hypoxia development and continuation in many areas of the world's coastal ocean is accelerated by human activities, especially where nutrient loading increased in the Anthropocene. This higher loading set in motion a cascading set of events related to eutrophication. The formation of hypoxic areas has been exacerbated by any combination of interactions that increase primary production and accumulation of organic carbon leading to increased respiratory demand for oxygen below a seasonal or permanent pycnocline. Nutrient loading is likely to increase further as population growth and resource intensification rises, especially with increased dependency on crops using fertilizers, burning of fossil fuels, urbanization, and waste water generation. It is likely that the occurrence and persistence of hypoxia will be even more widespread and have more impacts than presently observed. Global climate change will further complicate the causative factors in both natural and human-caused hypoxia. The likelihood of strengthened stratification alone, from increased surface water temperature as the global climate warms, is sufficient to worsen hypoxia where it currently exists and facilitate its formation in additional waters. Increased precipitation that increases freshwater discharge and flux of nutrients will result in increased primary production in the receiving waters up to a point. The interplay of increased nutrients and stratification where they occur will aggravate and accelerate hypoxia. Changes in wind fields may expand oxygen minimum zones onto more continental shelf areas. On the other hand, not all regions will experience increased precipitation, some oceanic water temperatures may decrease as currents shift, and frequency and severity of tropical storms may increase and temporarily disrupt hypoxia more often. The consequences of global warming and climate change are effectively uncontrollable at least in the near term. On the other hand, the consequences of eutrophication-induced hypoxia can be reversed if long-term, broad-scale, and persistent efforts to reduce substantial nutrient loads are developed and implemented. In the face of globally expanding hypoxia, there is a need for water and resource managers to act now to reduce nutrient loads to maintain, at least, the current status.

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Dynamics and distribution of natural and human-caused hypoxia

Denitrification, the reduction of the nitrogen (N) oxides, nitrate (NO3-) and nitrite (NO2-), to the gases nitric oxide (NO), nitrous oxide (N2O), and dinitrogen (N2), is important to primary production, water quality, and the chemistry and physics of the atmosphere at ecosystem, landscape, regional, and global scales. Unfortunately, this process is very difficult to measure, and existing methods are problematic for different reasons in different places at different times. In this paper, we review the major approaches that have been taken to measure denitrification in terrestrial and aquatic environments and discuss the strengths, weaknesses, and future prospects for the different methods. Methodological approaches covered include (1) acetylene-based methods, (2) 15N tracers, (3) direct N2 quantification, (4) N2:Ar ratio quantification, (5) mass balance approaches, (6) stoichiometric approaches, (7) methods based on stable isotopes, (8) in situ gradients with atmospheric environmental tracers, and (9) molecular approaches. Our review makes it clear that the prospects for improved quantification of denitrification vary greatly in different environments and at different scales. While current methodology allows for the production of accurate estimates of denitrification at scales relevant to water and air quality and ecosystem fertility questions in some systems (e.g., aquatic sediments, well-defined aquifers), methodology for other systems, especially upland terrestrial areas, still needs development. Comparison of mass balance and stoichiometric approaches that constrain estimates of denitrification at large scales with point measurements (made using multiple methods), in multiple systems, is likely to propel more improvement in denitrification methods over the next few years.

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Methods for measuring denitrification: diverse approaches to a difficult problem

The osmoadaptation of most micro-organisms involves the accumulation of K+ ions and one or more of a restricted range of low molecular mass organic solutes, collectively termed ‘compatible solutes’. These solutes are accumulated to high intracellular concentrations, in order to balance the osmotic pressure of the growth medium and maintain cell turgor pressure, which provides the driving force for cell extension growth. In this review, I discuss the alternative roles which compatible solutes may also play as intracellular reserves of carbon, energy and nitrogen, and as more general stress metabolites involved in protection of cells against other environmental stresses including heat, desiccation and freezing. Thus, the evolutionary selection for the accumulation of a specific compatible solute may not depend solely upon its function during osmoadaptation, but also upon the secondary benefits its accumulation provides, such as increased tolerance of other environmental stresses prevalent in the organism’s niche or even anti-herbivory or dispersal functions in the case of dimethylsulfoniopropionate (DMSP). In the second part of the review, I discuss the ecological consequences of the release of compatible solutes to the environment, where they can provide sources of compatible solutes, carbon, nitrogen and energy for other members of the micro-flora. Finally, at the global scale the metabolism of specific compatible solutes (betaines and DMSP) in brackish water, marine and hypersaline environments may influence global climate, due to the production of the trace gases, methane and dimethylsulfide (DMS) and in the case of DMS, also couple the marine and terrestrial sulfur cycles.

Ecological significance of compatible solute accumulation by micro‐organisms: from single cells to global climate

Urbanization is a major cause of loss of coastal wetlands. Urbanization also exerts significant influences on the structure and function of coastal wetlands, mainly through modifying the hydrological and sedimentation regimes, and the dynamics of nutrients and chemical pollutants. Natural coastal wetlands are characterized by a hydrological regime comprising concentrated flow to estuarine and coastal areas during flood events, and diffused discharge into groundwater and waterways during the non-flood periods. Urbanization, through increasing the amount of impervious areas in the catchment, results in a replacement of this regime by concentrating rain run-off. Quality of run-off is also modified in urban areas, as loadings of sediment, nutrients and pollutants are increased in urban areas. While the effects of such modifications on the biota and the physical environment have been relatively well studied, there is to date little information on their impact at the ecosystem level. Methodological issues, such as a lack of sufficient replication at the whole-habitat level, the lack of suitable indices of urbanization and tools for assessing hydrological connectivity, have to be overcome to allow the effects of urbanization to be assessed at the ecosystem level. A functional model is presented to demonstrate the impact of urbanization on coastal wetland structure and function.

Impact of urbanization on coastal wetland structure and function

Nitrogen fixation in seagrass meadows: Regulation, plant–bacteria interactions and significance to primary productivity

Due to their boom and bust population dynamics and the enormous biomasses they can attain, jellyfish and ctenophores can have a large influence on the cycling of carbon (C), nitrogen (N) and phosphorus (P). This review initially summarises the biochemical composition of jellyfish, and compares and contrasts the mechanisms by which non-zooxanthellate and zooxanthellate jellyfish acquire and recycle C, N and P. The potential influence of elemental cycling by populations of jellyfish on phytoplankton and bacterioplankton production is then assessed. Non-zooxanthellate jellyfish acquire C, N and P predominantly through predation on zooplankton with smaller contributions from the uptake of dissolved organic matter. C, N and P are regenerated via excretion of inorganic (predominantly ammonium (NH4
 +) and phosphate (PO4
 3−)) and dissolved organic forms (e.g. dissolved free amino acids and dissolved primary amines). Inorganic nutrients excreted by jellyfish populations provide a small but significant proportion of the N and P required for primary production by phytoplankton. Excretion of dissolved organic matter may also support bacterioplankton production but few data are available. In contrast, zooxanthellate medusae derive most of their C from the translocation of photosynthetic products, exhibit no or minimal net release of N and P, and may actively compete with phytoplankton for dissolved inorganic nutrients. Decomposition of jellyfish blooms could result in a large release of inorganic and organic nutrients and the oxygen demand required to decompose their tissues could lead to localised hypoxic or anoxic conditions.

Influence of jellyfish blooms on carbon, nitrogen and phosphorus cycling and plankton production

Ammonium is an important nutrient in primary production; however, high ammonium loads can cause eutrophication of natural waterways, contributing to undesirable changes in water quality and ecosystem structure. While ammonium pollution comes from diffuse agricultural sources, making control difficult, industrial or municipal point sources such as wastewater treatment plants also contribute significantly to overall ammonium pollution. These latter sources can be targeted more readily to control ammonium release into water systems. To assist policy makers and researchers in understanding the diversity of treatment options and the best option for their circumstance, this paper produces a comprehensive review of existing treatment options for ammonium removal with a particular focus on those technologies which offer the highest rates of removal and cost-effectiveness. Ion exchange and adsorption material methods are simple to apply, cost-effective, environmentally friendly technologies which are quite efficient at removing ammonium from treated water. The review presents a list of adsorbents from the literature, their adsorption capacities and other parameters needed for ammonium removal. Further, the preparation of adsorbents with high ammonium removal capacities and new adsorbents is discussed in the context of their relative cost, removal efficiencies, and limitations. Efficient, cost-effective, and environmental friendly adsorbents for the removal of ammonium on a large scale for commercial or water treatment plants are provided. In addition, future perspectives on removing ammonium using adsorbents are presented.

David T. Welsh

Papers

Ecological significance of compatible solute accumulation by micro‐organisms: from single cells to global climate

Impact of urbanization on coastal wetland structure and function

Nitrogen fixation in seagrass meadows: Regulation, plant–bacteria interactions and significance to primary productivity

Influence of jellyfish blooms on carbon, nitrogen and phosphorus cycling and plankton production

Removing ammonium from water and wastewater using cost-effective adsorbents: A review.